TEXAS TECH UNIVERSITY
Natural Science Research Laboratory
Special Publications
Museum of Texas Tech University
Number 53 15 February 2008
Nocturnal Lemur Diversity at
Masoala National Park
Moore’s YVooJly Lemur
Avahi mooreorum
Scott's Sportive Lemur
Lepifemur scoftoruni
40 Km
Masoala Peninsula
Masiaposa Forest
Runhua Lei, Shannon E. Engberg, Rambinintsoa Andriantompoha van a, Susie M. McGuire,
Russell A. Mittermeier, John R. Zaonarivelo, Rick A. Brenneman, and Edward E. Louis, Jr.
Front cover: Distribution of two new species of nocturnal lemurs at Masoala National Park, Madagascar. Descriptions
of Moore’s woolly lemur, Avahi mooreorum, and Scott’s sportive lemur, Lepilemur scottorum, are presented in this
publication. Figure created by Shannon E, Engberg, Rick A. Brenneman, and Edward E. Louis, Jr.
Special Publications
Museum of Texas Tech University
Number 53
Nocturnal Lemur Diversity
at Masoala National Park
RunhuaLei, Shannon E. Engberg, Rambinintsoa Andriantompohavana, Susie M. McGuire,
Russell A. Mittermeier, John R. Zaonarivelo, Rick A. Brenneman, and Edward E. Louis, Jr.
Henry Doorly Zoo, Conservation International, and
University of Antananarivo
Layout and Design: Jacquel ine Chavez and Lisa Bradley
Cover Design: Shannon E, Engberg, Rick A. Brenneman, and Edward E. Louis, Jr.
Copyright 2008, Museum of Texas Tech University
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Printed: 15 February 2008
Library of Congress Cataloging-in-Publication Data
Special Publications of the Museum of Texas Tech University, Number 53
Series Editor: Robert J. Baker
Nocturnal Lemur Diversity at Masoala National Park
Runhua Lei, Shannon E. Engberg, Rambinintsoa Andriantompohavana, Susie M. McGuire, Russell A.
Mittermeier, John R. Zaonarivelo, Rick A. Brenneman, and Edward E. Louis, Jr.
ISSN 0169-0237
ISBN 1-929330-15-4
ISBN13 978-1-929330-15-7
Museum of Texas Tech University
Lubbock, TX 79409-3191 USA
(806)742-2442
Nocturnal Lemur Diversity at Masoala National Park
RunhuaLei, Shannon E. Engberg, Rambinintsoa Andriantompohavana, Susie M. McGuire, Russell A.
Mittermeier, John R. Zaonarivelo, Rick A. Brenneman, and Edward E. Louis, Jr.
Abstract
Approximately 3,000 base pairs of mitochondrial DNA sequence data were
utilized as part of a phylogenetic analysis to investigate the taxonomic relationships
of the nocturnal lemurs (genera Avahi and Lepilemur) of Masoala National Park. The
molecular data distinguished all currently described Avahi and Lepilemur species, but
it also revealed previously unrecognized biodiversity within these genera at Masoala
National Park. Diagnoses of a new species of woolly lemur and a new species of sportive
lemur from Masoala National Park are presented. These results demonstrate that cryptic
species can be revealed through molecular data and the importance of initiating further
detailed studies in previously unstudied sites to confirm the distributions of all species,
particularly nocturnal lemur species.
Key words: Avahi , D-loop, Lepilemur , Madagascar, Masoala National Park,
Pastorini fragment
Introduction
Madagascar, with a land mass of 587,000 km 2
spanning from latitudes 12°S to 25°S, has a diverse
range of habitats created by an interaction of variable
abiotic factors such as temperature and rainfall (north-
south gradient) and altitude (gradients that segregate
the island into generally eastern and western sections
(Donque 1972; Stevens and O’Connor 2006)). Based
on high levels of endemism and species diversity but
confronted by persistent human related pressures,
Madagascar is considered one of the most critical global
priorities for conservation protection and has been des¬
ignated as one of the world’s most important biodiver¬
sity hotspots (Green and Sussman 1990; Myers 2000;
Groombridge and Jenkins 2002). Prominently figured
into conservation efforts, the lemurs of Madagascar are
protected under the Convention of international Trade
of Endangered Species (CITES) and are designated
by the 1UCN/SSC Red List Categories from critically
endangered to threatened or as data deficient (1UCN
2004). Directed almost entirely towards nocturnal le¬
murs, recent molecular and morphological studies have
led to a significant increase in the number of recognized
species (Groves 2000; Rasoloarison et al. 2000; Thal-
mann and Geissmann 2000,2005; Kappeler et al. 2005;
Andriaholinirina et al. 2006; Andriantompohavana et
al. 2006,2007; Louis etal. 2006a, 2006b; Olivieri et al.
2007). Given the numerous taxonomic revisions and
newly described species that have been presented over
the past decade, a comprehensive understanding of the
distribution and taxonomy of the nocturnal lemurs is
needed. We contribute to this line of research by pre¬
senting data on the taxonomy of two nocturnal genera,
Lepilemur and Avahi, concentrating on the systematics
of these taxa at Masoala National Park.
The Masoala Peninsula, located in northeastern
Madagascar, contains one of the largest remaining tracts
of humid evergreen forest, including significant areas
of lowland forest (Fig 1; Kremen et al. 1999). During
an initial lemur survey of the peninsula. Sterling and
Rakotoarison (1998) identified nine species including
Lepilemur mustelinus and Avahi laniger. Conducting
rapid lemur surveys of Masoala National Park, Vasey
(2000) and Mittermeier et al. (2006) also identified nine
1
2
Special Publications, Museum of Texas Tech University
data from Andriantompohavana et al. (2007) and Louis et al. (2006b; Appendix 11(a)).
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
3
species of lemurs, including the same two nocturnal
lemurs. Ganzhorn et al. (2006), Louis et al. (2006b),
Craul et al. (2007), and Oliver! et al. (2007), illustrated
the importance of Madagascar’s rivers as barriers to
migration, promoting the processes of allopatric specia-
tion. The Antainambalana River, a prominent river to
the west of Masoala Peninsula, is known to be a bar¬
rier to several species such as Propithecus diadema to
Propithecus ccmdidus,. and Varecia variegata to Varecia
rubra (F ig. 1). Furthermore, Simpson (1964) described
the potential isolating effects of peninsulas on fauna
and flora, citing as an example the Florida peninsula.
On peninsulas, there is an inverse relationship between
the number of species and geographic distance from the
mainland, which acts as a reservoir of genetic diversity.
This potential isolating effect has been documented
with Varecia rubra in the Masoala Peninsula, but its
significance has not been demonstrated with other taxa.
Given the potential significance of rivers and peninsulas
on speeiation, the taxonomic status of the Lepilemur
and Avahi species should be established. Therefore, a
molecular analysis was conducted in order to clarify
the nocturnal lemur diversity of the genera Avahi and
Lepilemur at Masoala National Park.
The genus Avahi was originally described as a
single species, Avahi laniger, with two subspecies:
A. laniger occidentalis in northern, northwestern,
and western forests, and A. laniger laniger in the
eastern forests (Petter et al. 1977; Tattersall 1982).
Subsequently, based on cytogenetic studies, Rumpler
et al. (1990) elevated them to full species status as A.
occidentalis and A. laniger. According to morphologi¬
cal and vocalization data, Thalmann and Geissmann
(2000) described a third species, A. unicolor , from the
Ampasindava Peninsula, and more recently discovered
a fourth species, A. cleesei , from the Tsingy de Bema-
raha region (Thalmann and Geissmann 2005). Futher
revisions of this genus by Zaramody et al. (2006) and
Andriantompohavana et al. (2007) expanded the num¬
ber of recognized species in the genus Avahi to eight full
species. Based on the distribution of woolly lemurs,
the Avahi species found in the Masoala Peninsula has
been understood to be the eastern woolly lemur, Avahi
laniger.
The sportive lemurs, genus Lepilemur , are me¬
dium-sized, nocturnal lemurs found in virtually all
forested areas in Madagascar (Mittermeier et al. 2006).
Initially, Schwarz (1931) and Hill (1953) recognized
two species of Lepilemur. L. mustelinus from the east¬
ern rainforests and L. ruficaudatus from the western and
southern dry forests of Madagascar. L. ruficaudatus
was further divided into two subspecies, L. ruficaudatus
ruficaudatus and L. ruficaudatus leucopus. Since then,
the taxonomy of the sportive lemurs has been revised
repeatedly (Petter and Petter-Rousseaux 1960; Rumpler
and Albignac 1975; Petter et al. 1977; Tattersall 1982;
Jenkins 1987; Mittermeier et al. 1994; Ravoarima-
nana et al. 1999, 2004; Thalmann 2000; Groves 2001;
Rumpler et al. 2001; Thalmann and Ganzhorn 2003).
More recently, the genus Lepilemur has undergone a
dramatic expansion to 24 recognized species with the
taxonomic revisions of Andriaholinrrina et al. (2006;
described three new species), Louis et al. (2006b;
described 11 new species), Rabarivola et al. (2006;
described one new species), and Craul et al. 2007;
described two new species). Additionally, Zinner et
al. (2007) raised important concerns with the sportive
lemurs recently described in northwestern Madagascar,
citing the need for comparative analyses including the
holotype specimens (specifically L. dorsalis and L.
grandidieri) and respective data sets, along with an
indepth morphological analysis. Based on the distribu¬
tion of sportive lemurs, the Lepilemur species found in
the Masoala Peninsula has been understood to be the
weasel sportive lemur, Lepilemur mustelinus.
Historically, sympatric reproductive isolation,
described in the Biological Species Concept (BSC),
predominantly was used as criterion defining species
(Mayr 1942). However, when the putative species is
a geographically isolated or an allopatrically defined
population, this concept is difficult to implement. The
Phylogenetic Species Concept (PSC) not only en¬
compasses a cladistic perspective, but also allows the
scientist to focus operationally on the results of evolu¬
tion to present and to delineate species (Cracraft 1983;
Wheeler and Platnick 2000; Louis et al. 2006a).
In this paper, we present comparative phyloge¬
netic analyses of mitochondrial DNA sequence, aug¬
mented with morphological and phenotypic character
data, of the nocturnal lemurs from the genera Avahi
and Lepilemur at Masoala National Park to recently
recognized woolly (Andriantompohavana et al. 2007)
and sportive lemurs (Louis et al. 2006b). The results
of phylogenetic analyses of molecular sequence data
4
Special Publications, Museum of Texas Tech University
for the genera Lepilemur and Avahi reveal unique bio¬
diversity at Masoala National Park, Madagascar. Diag¬
nostic descriptions of this previously undescribed lemur
diversity are presented. Additionally, an electronic
database that includes all Avahi and Lepilemur field
data and photographs, including data for the paratype
specimens, is curated at the Museum of Texas Tech Uni¬
versity. The database is stored in the Type Specimen
Collection in multiple media formats. This collection of
field data and photographs, as well as additional tables
and figures, also are available online at the website of
Omaha's Henry Doorly Zoo. See Appendices I-II1 for
a directory of appropriate website addresses.
Methods
Sampling. —All lemurs investigated in this study
were wild-caught and were immobilized with a CO,
projection rifle or blowgun with 10 mg/kg of Telazol
(Fort Dodge Animal Health; Overland Park, KS; Ap¬
pendix I(a-b)). All recently recognized woolly and
sportive lemurs were included in the analysis except
for L. mittermeieri (Rabarivola et al. 2006). Rabarivola
et al. (2006) utilized cytogenetic and cytochrome b
subunit sequence data in their analyses, and since no
sample was available to generate comparative sequence
data, we could not include this species in this study.
Four 2.0 mm biopsies and 1.0 cc per kilogram of whole
blood were collected from each sedated animal and
immediately stored in room temperature storage buf¬
fer (Longmire et al. 1992). We recorded the location
of the immobilized lemurs using a global positioning
system (Appendix I(a-b)). Beginning in 2000, while
the animal was sedated, we placed a HomeAgain
(Schering-Plough Veterinary Corp.; Kenilworth, NJ)
microchip subcutaneously between the scapulae of
each lemur (Appendix I(a-b)). This procedure was
used to field catalog each animal with a unique recog¬
nition code to provide for the capability to re-identify
positively all captured individuals during any future
immobilizations.
Measurements were taken on the sedated animals
as described in Andriantompohavana et al. (2007) and
Louis et al. (2006b). For presentation purposes, we
present the weight, head crown, body length, and tail
length in this publication following the guidelines of
Smith and Jungers (1997; Appendix 1(a)). All other
measurements, e-voucher photographs, and data are
available at http://www.omahazoo.com/ccr/index.
asp?page=/ccr/genetics/genhome.htm. All traits were
tested for significance among species using ANalysis
Of VAriance (ANOVA) performed using an online
statistical service (Kirkman 1996). Species means
were plotted in histogram form with the appropriate
95% confidence intervals.
As previously discussed in Thalmann and Geiss-
mann (2005), Andriantompohavana etal. (2006), Louis
et al. (2006a, 2006b), and Andriantompohavana et al.
(2007), the utilization of whole vouchers as the desig¬
nated holotype for a new species is not a prerequisite;
however, opportunistic collections can later supplement
morphological and/or molecular data in combination
with curated blood and/or tissue samples. The woolly
and sportive lemurs are prime candidates for this meth¬
odology since the highly folivorous dietary require¬
ments of these groups of lemurs currently precludes
any attempts to curate ‘live vouchers” (Thalmann and
Geissmann 2005; Louis et al. 2006b; Andriantompo¬
havana et al. 2007). Total genomic DNA for the three
paratype specimens of Avahi and Lepilemur species
are currently curated at the Museum of Texas Tech
University (TK145356; TK145357; TK145358; and
TK145333; TK145334; TK145335, respectively).
Data Collection. —Genomic DNA was extracted
from a 2.0 millimeter ear punch using phenol-chloro¬
form extraction (Sambrook et al. 1989; Appendix 1(b)).
To correlate our data with previously published molecu¬
lar studies, we analyzed the following regions of the
mitochondrial DNA(mtDNA): the displacement loop
or control region (D-loop; Baker et al. 1993; Wyner et
al. 1999); the 12S subunit ribosomal RNA gene, along
with a portion of the tRNA Phe (only the Lepilemur indi¬
viduals; 12S rRNA; Hedges 1994; Wyner etal. 1999);
and a fragment of the cytochrome oxidase subunit III
gene (COIII), NADH-dehydrogenase subunits 3, 4L,
and 4 (ND3, ND4L, and ND4), as well as the tRNA Gly ,
tRNA Arg , tRNA His , tRNA Ser , and partial tRNA Leu genes
(subsequently referred to as the PAST fragment; Pas-
torini et al. 2000). Using 50 nanograms of genomic
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
5
DNA, the D-loop and the PAST (2386 bp) fragments
were amplified using the following conditions: 94°C
for 30s, 47°C for 45s, 72°C for 45s for 34 cycles.
Since potential nuclear insertions or mitochondrial
pseudogenes within the nuclear genome can be ampli¬
fied inadvertently, we chose to minimize this likeli¬
hood by amplifying both mitochondrial DNA regions
as intersecting or overlapping segments (Zhang and
Hewitt 1996). Consequently, the PAST fragment was
generated from five amplified segments. Additionally,
to eliminate further amplification of nuclear insertions,
a rapid, efficient, and species independent technique
derived from the degenerate oligonucleotide-primed
PCR method (DOP-PCR) was used to generate the PCR
products (Teleniusetal. 1992). Adapting this LL-DOP-
PCR (long products from low quantity), the sequence
data were generated from overlapping segments to con¬
firm the concensus D-loop and PAST fragments. The
samples were electrophoresed in a 1 .2% agarose gel to
verify the PCR product and purified using QIAquick
PCR purification kit (Q1AGEN; Valencia, CA). The
cleaned products were cycle sequenced using a big dye-
terminator sequencing kit (Applied Biosystems; Foster
City, CA). The sequences were analyzed by capillary
electrophoresis with an Applied Biosystems Prizm 3100
genetic analyzer. A suite of internal sequencing primers
from Pastorini et al. (2000) and Pastorini et al. (2001)
were used to generate the PAST fragment. Addition¬
ally, PCR and sequencing primers specific for Avahi
and Lepilemur were designed for the PAST and D-loop
fragments (Louis et al. 2006b; Andriantompohavana
et al. 2007). The sequence fragments were aligned to
generate a consensus sequence using Sequencher (Gene
Corp; Ann Arbor, Ml), and the consensus sequences
were aligned using Clustal X (Thompson et al. 1997).
All sequences have been deposited in GenBank and
the sequence data and information are available from
the referenced accession numbers (Appendix I(a-b)).
The sequence alignments for the data sets are available
from the first author upon request.
Phylogenetic A na lysis .—Maxi mum- pars i mo ny
(MP), maximum likelihood (ML), and neighbor-join-
ing (NJ) analyses were performed for the phylogenetic
study of the D-loop, PAST, 12S rRNA (Lepilemur
only), and combined fragments (D-loop and PAST
fragments for Avahi and Lepilemur) sequence data with
PAUP* 4.0b 10 software (Swofford 2001). The trees
described in this paper are all consensus trees except
for the bootstrap analysis (all trees were presented as
phylograms for presentation purposes only). Bootstrap
analyses were accomplished with 1000, 1000, 3000,
and 4000 replicates with the D-loop, 12S rRNA, PAST,
and D-loop /PAST sequence files, respectively, with 10
random addition heuristic searches per replicate option
selected. Only nodes with greater than 50% support
were reported. The D-loop NJ tree was generated
using the Tamura-Nei model (Tamura and Nei 1993).
The stepwise addition option was selected for MP and
ML analyses, and corrections for nucleotide sequence
data suggested by Kimura (1980) were used with the
NJ analyses. Gaps were considered as a fifth character
in MP analyses, whereas gaps were treated as missing
data in the NJ analyses. The ML trees were estimated
via the heuristic search. For the substitution model,
the transition/transversion ratios were estimated in
MacCIade 3.01 (Maddison and Maddison 1992) and
a discrete approximation to gamma distribution was
estimated for among site rate variation. The default
settings were maintained for all other settings, thus
yielding the equivalent of the HKY model (Hasegawa
et al. 1985). In addition to character-based phylogenetic
analysis of DNA sequences, PAUP* 4.0bl0 software
(Swofford 2001) and MEGA 3.1 (Kumar et al. 2004)
were used to calculate uncorrected pairwise distances
(‘p’) and Kimura distance measures for D-loop and
PAST fragments.
Bayesian inference analyses were conducted
using MrBayes 3.0b4 (Huelsenbeck and Ronquist
2001; Ronquist and Huelsenbeck 2003). The model
of evolution was selected by using Mrmodeltest 2.2, a
modified version of Modeltest 3.6 (Posada and Cran¬
dall 1998; Ny lander 2004). A Markov Chain Monte
Carlo (MCMC) run with four simultaneous chains
and 1,000,000 generations was performed. Every
hundredth generation, the tree with the best likelihood
score was saved, resulting in 4,000 trees. The 4,000
trees were condensed in a majority rule consensus
tree using PAUP* 4.0b 10 (Swofford 2001). Branch
supports were assigned as posterior probabilities on
the consensus tree. The pattern of sequence evolution
was estimated by conducting a minimum spanning
network generated with the program NETWORK 4.11
(Bandelt et al. 1999; Forster et al. 2001) and Arlequin
2.0 (Schneider et al. 2000).
As described in Davis and Nixon (1992), Wyner
et al. (1999), Mayor et al. (2004), Andriantompohavana
et al. (2006), Louis et al. (2006a, 2006b), and Andrian-
6
Special Publications, Museum of Texas Tech University
tompohavana et al. (2007), we utilized MacClade 3.01
(Maddison and Maddison 1992) and MEGA3.1 (Ku¬
mar et al. 2004) to designate evolutionary significant
units (ESU) for the genera Avahi and Lepilemur using
a population aggregate analysis (PAA) of the D-loop
and PAST sequence data. With the sequential addition
of each individual without an a priori species desig¬
nation, a PAA distinguishes attributes or apomorphic
characters according to the smallest definable unit
(Davis and Nixon 1992; Ravaoarimanana et al. 2004;
Mayor et al. 2004; Andnantompohavana et al. 2006;
Louis et al. 2006a, 2006b; Andnantompohavana et al.
2007; Olivieri et al. 2007).
Results
Population Aggregate Analysis for the genera
Avahi and Lepilemur.—A summary of the PAA for the
diagnostic s ites for the genera Avahi and Lepilemur are
condensed in Tables 1A-B and Tables 1C-E, respec¬
tively, The results from the PAA of the D-loop and
PAST sequence data for the genus Avahi are presented
as diagnostic nucleotide sites in Appendices lll(a-b),
respectively. The results from the PAA of the D-loop,
12 S rRNA, and PAST sequence data for the genus
Lepilemur are presented as diagnostic nucleotide sites
in Appendices Ill(c-e), respectively. Multiple diagnos¬
tic characters differentiated each Avahi species, along
with two types of A. laniger (only the results from the
PAST fragment PAA designated attributes for the two
types), three types of A. peyrierasi , and previously
unrecognized biodiversity, Avahi species nova #4 from
Masoala National Park (Tables 1A-B). Furthermore,
multiple diagnostic characters differentiated each
recognized Lepilemur , along with distinguishing two
distinct clades designated as Lepilemur species nova
#1 at Masoala National Park and Lepilemur species
nova #2 from Mananara-Nord National Park (Tables
1C-E).
Morphological Data of the genera Avahi and
Lepilemur.—Means and standard deviations as well
as P-values for significant differences among species
are presented in Table 2A-C for the Avahi species and
Table 3 A-C for the Lepilemur species. The histograms,
depicting the mean and 95% confidence interval of sig¬
nificant tests detecting morphometrical trends among
the species within a genus, and those morphological
units in which significant differences were identified
are presented {Avahi species nova #4, Lepilemur spe¬
cies nova # 1, and Lepilemur species nova #2 were
analyzed as separate taxa; Appendiix ll(a-d)). The raw
morphological and field data for woolly and sportive
lemur species are presented in Appendix 1(a).
Phylogenetic Analysis of the genus Avahi.—Mi¬
tochondrial DNA sequence data were completed for
two fragments, D-loop and PAST (approximately
3,000 bp) for 119 individuals, representing all eight
recognized species of woolly lemurs from a total of 14
sites (Appendices I(a-b); Andnantompohavana et al.
2007). Based on the phylogenetic reconstructions of
the NJ, MP, and ML analyses of three sequence align¬
ments (D-loop, PAST, and D-loop-PAST combined),
the eight recognized Avahi species are differentiated
and could be delineated into two major terminal clades
or subgroups (Fig. 2; Appendices ll(e-i)). The first
subgroup corresponds to the woolly lemurs of western
Madagascar as follows: A. occidentalism, cleesei, and
A. unicolor (Fig. 2). The second subgroup corresponds
to the eastern woolly lemurs as follows: A. laniger ,
A. meridionalis , A. ramanantsoavancr A . peyrierasi ,
and A. betsileo (Fig. 2). Three haplotype groups or
types of A. peyrierasi were distinguishable within the
woolly lemurs at Ranomafana National Park (Fig. 2;
Appendices IT(e-i)). In addition, another group of
woolly lemurs from Masoala National Park, A. spe¬
cies nova #4, formed an apparent clade from the other
species (Fig. 2; Appendices IT(e-i)). High bootstrap
values support the MP, NJ, and Bayesian analyses with
respect to the topology of the genera and species (Fig.
2; Appendices II(f, h, and i)). The minimum spanning
network presents diagrammatically the speciation
among eight woolly lemurs, along with A. species
nova #4 (Fig. 3).
The uncorrected ‘p’ distance and the absolute
genetic differences for D-loop and PAST fragments
for the genus Avahi are presented in Appendix Ill(f-g).
The highest average uncorrected pairwise distances and
absolute genetic differences for the D-loop sequence
alignment were between the western and eastern woolly
(text continued on page 19)
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
7
Table l A. Summary of Population Aggregate Analysis (PAA) D-Loop fragment diagnostic sites for Avahi. Refer to Appendix III,
Table 1A. *No character or attribute is available for this fragment.
Species
Fragment Size
(bp)
PAA base pair location
Avahi peyrierasi type #2
562
172, 194, 195, 333
Avahi peyrierasi type #3
562
336, 548
Avahi peyrierasi type # 1
562
349
Avahi betsileo
562
20, 227, 277, 309, 521
Avahi meridionalis
562
16, 64, 159, 422, 506
Avahi ramanantsoavana
562
158, 194, 199, 525
Avahi laniger type #1 (Mantadia)
562
543
Avahi laniger type #2 (Mananara-Nord)
562
Avahi species nova #4
562
154, 331
Avahi unicolor
563
106,288,397, 401
Avahi cleesei
563
64, 275, 276, 287, 300, 302, 332
Avahi occidentalis
563
295, 296,314, 499
Table IB. Summary of Population Aggregate Analysis (PAA) PASTfragment diagnostic sitesfor Avahi. Refer to Appendix III, Table
IB. The locality q/Avahi laniger type #7 is Mantadia National Park. The locality q/Avahi laniger type #2 is Mananara-Nord National
Park. The locality q/Avahi species nova #4 is Masoala National Park. *No character or attribute is available for this fragment.
Fragment Size
Species
(bp)
PAA base pair location
Avahi peyrierasi type #2
2364
14, 57, 155, 344, 572,698,945, 1124, 1139, 1341, 1353, 1525,
1537, 1818, 1835,1886, 1960, 2059, 2111, 2122
Avahi peyrierasi type #3
2364
54, 246, 701, 1408, 1584, 1861, 1946, 2096, 2153
Avahi peyrierasi type #1
2364
620, 683, 1426, 1783,2165
Avahi betsileo
2364
920, 1053,1837
Avahi meridionalis
2364
85, 289, 291, 339, 708, 847, 935, 952, 958, 1060, 1114, 1131, 1270,
1336, 1339, 1459, 1612, 1771, 2013, 2356
Avahi ramanantsoavana
2364
117,279, 938, 940, 972, 985, 1261, 1273, 1381, 1384, 1407, 1807
Avahi laniger type #1
2364
671, 957,. 1117
Avahi laniger type #2
2364
576, 835
Avahi species nova #4
2364
288, 650, 844, 957, 997, 1009, 1114, 1141, 1157, 1183, 1288, 1362,
1597, 1675, 1729, 1846, 1933, 1963, 1996, 2011, 2029, 2101, 2141,
2170,2187,2210
Avahi unicolor
2364
6, 93, 114, 133, 501, 636, 916, 1052, 1135, 1279, 1454, 1849, 1872,
2032,2116,2131
Avahi cleesei
2364
8, 118, 213, 258, 522, 704, 928, 1057, 1213, 1273, 1552, 1585,
1786
Avahi occidentalis
2364
47, 111, 121, 500, 540, 665, 764, 783, 826, 1075, 1138, 1168, 1333,
1477, 1519, 1525, 2023, 2053, 2116, 2138, 2332
8
Special Publications, Museum of Texas Tech University
Table 1C. Summary of Population Aggregate Analysis (PAA) D-Loop diagnostic sites for Avahi. Refer to Appendix 111 , Table 1C.
The locality o/Lepilemur species nova #1 is Masoala National Park. The locality o/Uepilemur species nova #2 is Mananara-Nord
National Park. *No character or attribute is available for this fragment.
Species
Fragment Size
(bp)
PAA base pair location
Lepilemur ankaranensis
540
*
Lepilemur milanoii
540
130
Lepilemur tymerlachsoni
538
117
Lepilemur septentrional is
536
33, 37, 98, 111,249
Lepilemur dorsalis
540
536, 537
Lepilemur sahamalazensis
542
*
Lepilemur petteri
534
*
Lepilemur leucopus
535
19
Lepilemur ruficaudatus
535
103, 126, 249, 310, 535
Lepilemur hubbardorum
535
242, 253, 270, 302
Lepilemur randrianasoli
538
33, 272
Lepilemur edwardsi
545-546
127, 139, 538
Lepilemur grewcockorum
544
139, 195,357
Lepilemur ahmansonorum
542
*
Lepilemur aeeclis
537-538
21
Lepilemur mustelinus
552-553
*
Lepilemur jamesorum
552
132
Lepilemur betsileo
553
272, 273, 286
Lepilemur fleuretae
550
10, 24, 37, 287, 288, 314, 317, 330
Lepilemur microdon
530
25, 34, 107, 110, 120, 121, 123, 124, 125, 137, 139, 396, 554
Lepilemur wrightae
551
55,58,275, 301,476, 493
Lepilemur seali
550
54, 159, 221
Lepilemur species nova # 1
550
87, 160, 195, 231, 327, 475
Lepilemur species nova #2
550
24, 30, 140, 187, 266
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
9
Table ID. Summary ofPopulation Aggregate Analysis (PAA) 12S rRNA diagnostic sites for Lepilemur. Refer to Appendix III, Table
ID. The locality o/Lepilemur species nova #1 is Masoala National Park. The locality o/Lepilemur species nova #2 is Mananara-
Nord National Park. *No character or attribute is available for this fragment.
Species
Fragment Size
(bp)
PAA base pair location
Lepilemur ankaranensis
855
*
Lepilemur milanoii
855
498
Lepilemur tymerlachsoni
855
394, 693, 854
Lepilemur septentrional is
855
18,60, 110, 154, 334, 488
Lepilemur dorsalis
855
761,796
Lepilemur sahamalazensis
855
294, 766
Lepilemur petteri
855
240
Lepilemur leucopus
855
*
Lepilemur ruficaudatus
855
433
Lepilemur hubbardurom
855
340
Lepilemur randrianasoli
855
6, 242, 369, 391,418, 666
Lepilemur edwardsi
854
*
Lepilemur grewcockorum
854
*
Lepilemur ahmansonorum
855
413,641,723, 768
Lepilemur aeeclis
855
389, 556
Lepilemur mustelinus
854
*
Lepilemur jamesorum
854
66
Lepilemur betsileo
853
432
Lepilemur fleuretae
854
53, 500, 757
Lepilemur microdon
855
235, 270, 757, 786
Lepilemur wrightae
854
349, 746
Lepilemur seali
855
186, 322, 544
Lepilemur species nova # 1
855
245, 326
Lepilemur species nova #2
855
126, 241,799
10
Special Publications, Museum of Texas Tech University
Table IE. Summary of Population Aggregate Analysis (PAA) PASTfragment diagnostic sites for Lepilemur. Refer to Appendix
III, Table IE. The locality o/Lepilemur species nova #1 is Masoala National Park. The locality o/Lepilemur species nova #2 is
Mananara-Nord National Park. *No character or attribute is available for this fragment.
Species
Fragment Size
(bp)
PAA base pair location
Lepilemur ankaranensis
2359-2360
364, 858, 1315, 1804
Lepilemur milanoii
2359
342, 769, 1896
Lepilemur tymerlachsoni
2361
152, 1309, 1378, 1861, 1898, 1995
Lepilemur septentrionalis
2361
44, 113, 211,214,274, 353, 354, 533, 551, 555, 576, 674, 734,
1103, 1174, 1231, 1347, 1399, 1448, 1492, 1550, 1582, 1603,
1630,1777,2144,2146,2363
Lepilemur dorsalis
2360
579, 717, 746,1525,1780, 2163,2168, 2177, 2236
Lepilemur sahamalazensis
2360
204, 399
Lepilemur petteri
2360-2361
337, 578, 779, 957, 1615
Lepilemur leucopus
2360
220,448,719, 836, 1960
Lepilemur ruficaudatus
2361
94, 127,365,776, 1074, 1370, 1783, 1835, 1867, 1921
Lepilemur hubbardorum
2360
350, 543, 566, 629, 681, 1012, 1015, 1240, 1396, 1559, 1906,
1907,2111
Lepilemur randrianasoli
2360
191, 699, 849, 923, 982, 1018, 1035, 1053, 1432, 1444, 1753,
1981, 1988, 2250, 2267
Lepilemur edwardsi
2360
1018, 1114, 1343, 1474, 1979
Lepilemur grewcockorum
2360
406, 888, 896, 988, 1114, 1226, 1354, 1537, 2230
Lepilemur ahmansonorum
2360
46, 304, 350,1096, 1097, 1402, 1818, 2141, 2170
Lepilemur aeeclis
2360
535, 548, 563, 581,975, 1357, 1368, 1423, 1442, 1990, 2089,
2107
Lepilemur mustelinus
2359-2360
85, 982
Lepilemur jamesorum
2360
2144
Lepilemur betsileo
2360
8, 1057
Lepilemur fleuretae
2360
29, 103,269, 358, 534, 546, 553, 664, 1124, 1574, 2013, 2023
Lepilemur microdon
2361
146, 510, 581, 596, 826, 829, 1171, 1369, 1954, 1991, 2077,
2164
Lepilemur wrightae
2360
133, 663, 691, 871, 907, 942, 1058, 1117, 1120, 1837, 1856,
1936, 2041, 2096, 2181, 2185, 2331
Lepilemur seali
2361
124, 147,290, 626, 665, 692, 722, 1302, 1313, 1371, 1679,
1875, 1879, 1969
Lepilemur species nova #1
2361
86, 567, 1157, 1337, 1435, 1483, 1606,2165
Lepilemur species nova #2
2361
72, 256, 871, 1033, 1112, 1167, 1237, 1336, 1538, 1902
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
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11 Avahi species and types. (Individual morphological data available online; see Appendix la).
12
Special Publications, Museum of Texas Tech University
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13
Table 3A. Morphometric data collectedfrom sedated Lepilemur individuals (Individual morphological data available in Ap¬
pendix la).
Species Name
Common Name
N
Weight
(kg)
Head
Crown
(cm)
Body
Length
(cm)
Tail
Length
(cm)
Lepilemur tymerlachsoni
Hawk’s sportive lemur
10
0.88±0.10
6.5±0.2
23.U1.5
24.7±2.2
Lepilemur aeeclis
Antafia sportive lemur
9
0.87±0.17
7.5±1.1
23.9±2.0
25.5d=0.9
Lepilemur ahmansonorum
Ahmanson’s sportive lemur
4
0.61±0.15
6.2±1.2
20.6±1.6
23.8±0.7
Lepilemur petteri
Petter’s sportive lemur
5
0.63±0.05
5.1±0.4
23.2±0.9
23.7±1.4
Lepilemur randrianasoli
Randrianasolo’s sportive
lemur
3
0.92±0.08
8.2±0.6
24.1±1.7
27.4±1.8
Lepilemur ankaranensis
Ankarana sportive lemur
30
0.78±0.12
7.0±0.8
21.8±1.9
27.7±2.1
Lepilemur septentrionalis
Northern sportive lemur
2
0.58±0.18
8.3±1.1
18.7±1.7
24.8±4.5
Lepilemur milanoii
Daraina or Swimming sportive
lemur
11
0.71±0.07
5.8±0.4
20.0±0.9
24.3±1.6
Lepilemur betsileo
Betsileo sportive lemur
2
1.10±0.15
7.5±0.4
25.3±0.1
27.2±1.2
Lepilemur dorsalis
Grey-Backed sportive lemur
6
0.75±0.07
7,5±0.4
23.2±1.5
27.5i0.8
Lepilemur edwardsi
Milne-Edwards’ sportive
lemur
11
U0i0.ll
6.7±0.6
26.3±2.7
28.4±2.1
Lepilemur fleuretae
Fleurete’s sportive lemur
1
0.80±0.00
7.1±0.0
25.8±0.0
29.0±0.0
Lepilemur sahamalazensis
Sahamalaza sportive lemur
2
0.70±0.07
6.3±0.1
19.6±0.3
23.7±2.7
Lepilemur grewcockorum
Grewcock’s sportive lemur
2
0.90±0.00
6.2±0.0
26.0±1.0
29.5±0.8
Lepilemur hubbardorum
Hubbard’s sportive lemur
7
0.99±0.18
7.6±1.4
24.1±0.8
24.0±1.0
Lepilemur ritftcaudatus
Red-tailed sportive lemur
3
0.86±0.06
6.9±1.0
24.3±1.7
24.2±3.6
Lepilemur jamesorum
James’ sportive lemur
8
1.05±0.08
8.0±0.3
27.1±2.7
30.0±1.7
Lepilemur leucopus
White-Footed sportive lemur
17
0.55±0.07
6.5±0.6
19.6±1.4
24.2±1.2
Lepilemur mustelinus
Weasel sportive lemur
27
0.99±0.21
7.9±1.4
25.8±4.0
25.2±2.1
Lepilemur wrightae
Wright’s sportive lemur
5
1.15±0.12
8.1±1.5
26.5±1.6
25.7±1.8
Lepilemur microdon
Small-Toothed sportive lemur
12
1.07±0.32
10.6±2.4
22.6±1.0
24.2±2.6
Lepilemur seali
Seal’s sportive lemur
5
0.95±0.09
7.5±0.9
27.5±1.4
26.0±1.4
Lepilemur species nova #1
sportive lemur new species # 1
5
0.88±0.14
6.7±0.6
26.6±0.9
27.8±2.0
Lepilemur species nova #2
sportive lemur new species #2
1
0.95±0.00
6.8±0.0
26.4±0.0
27.5±0.0
14 Special Publications, Museum of Texas Tech University
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Lepilemur Species P<0.0001 Weight Head Length Length Thumb Digit Hand Ulna merus Thumb Digit Foot Tibia Femur
L. mustelinus average 987.9 7.555 25.587 25.374 2.619 3.544 6.192 8.010 6.355 4.671 2.892 9.152 9.658 10.865
16
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Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
17
KANG45
—ANAL2.:
ANK7 — biicrucehas ravetobrma
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47
T*MIZA5.3
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—ANAL I &-- Pmpilhecta perrieri
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Artrifi Juritvcf
Avuhi rlfi^vi
m-.viAft
J-ALYJ
IMArM .4vtfAr tKcidunfalix
East Coast
Woolly Lemurs
West Coast
Woolly Lemurs
Figure 2. Maximum parsimony phylogram derived from the D-loop sequence data from 44 haplotypes
from the 119 Avahi individuals with 26 out-group taxa (one of 4031 most parsimonious trees). Values
above branches indicate number of changes between nodes. Values below branches indicate support
of bootstrap pseudoreplicates. Length = 1540; Cl = 0.5584; RI = 0.8618; RC = 0.4813; HI = 0.4416.
Special Publications, Museum of Texas Tech University
st
' y
1 — 3
.1 I
1 j c fct
4! 4
a. C
fc s a -a s s
e
■Ei ~ £ 'K V K
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Figure 3. Minimum spanning network of Avahi D-loop haplotypes calculated using Arlequin 2.0 and Network 4.11. Identification numbers denote haplotypes
corresponding to Appendix III(k). The minimum number of mutational steps separation matriarchal lines is indicated. Nucleotide substitutions are indicated
by dashes. The number of nucleotide differences (more than two) in their connecting lines of the network is indicated by the number at each connecting link.
Missing intermediates are indicated by conical little purple circles. The size of circles approximates the number of individuals with matching haplotypes
(circles without any number represent one individual).
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
19
lemurs, ranging from 12,2 to 8.6 and 66 to 50 (A. cleesei
to A. peyrierasi type #2, respectively). The highest
average uncorrected pairwise distances and absolute ge¬
netic differences for the PAST sequence alignment were
between the western and eastern woolly lemurs, ranging
from 13.2 to 10.2 and 283 to 220, respectively.
Phylogenetic Analysis of genus Lepilemur.—Mi¬
tochondrial DNA sequence data were completed for
three fragments, D-loop, 12S rRNA and PAST (ap¬
proximately 3,800 bp) for 216 individuals, representing
22 recognized species of sportive lemurs from a total
of 42 sites (Appendices I(a-b); Louis et al. 2006b).
Based on the phylogenetic inferences of the NJ, MP
and ML analyses of four sequence alignments (D-loop,
12S rRNA, PAST, and D-loop-PAST combined), the
22 recognized Lepilemur species are differentiated
and divided into five terminal clades (Fig. 4; Appendix
II(j-r)). The first terminal clade or subgroup consists
of sportive lemurs from northern Madagascar as fol¬
lows: L. ahmansonorum , L. ankaranensis , L. milanoii ,
L. septentrionalis ; L. tymerlachsoni , L. dorsalis, and L.
sahamalazensis (Fig. 4). The second terminal clade of
sportive lemurs is associated with southern Madagascar
and allied to the west-central subgroup (third terminal
clade) as follows: L. petteri and L. leucopus (Fig. 4).
The third terminal clade of sportive lemurs corresponds
to west-central Madagascar as follows: L. aeeclis , L.
randrianasoli, L. ruficaudatus , and L. huhbardorum
(Fig. 4). The fourth terminal clade of sportive lemurs
corresponds to eastern Madagascar as follows; L.
mustelinus , L. jamesorum , L. betsileo , L. fleuretae , L.
wrightae , L .seali , L. species nova #1, and L. species
nova #2 (Fig. 4). The fifth subgroup included two
western sportive lemurs and one eastern sportive lemur,
L. edwardsi and L. grewcockorum, and L. microdon ,
respectively (Fig. 4). There is high bootstrap support
for the MP, NJ, and Bayesian analyses with respect to
the topology of the genera and species (Fig. 4; Appen¬
dix II(j, 1, n, o, and r)).
The uncorrected ‘p’ distance and the absolute
genetic differences for the genus Lepilemur are pre¬
sented for D-loop, 12S rRNA, and PAST fragments
in Appendix lll(h-j), respectively. The highest and
lowest average uncorrected pairwise distances and
absolute genetic differences for the D-loop sequence
alignments between the sportive lemurs range from
18.6 to 2.7 and 107 to 20 (L. species nova #2 to L.
leucopus and L. ankaranensis to L. milanoii , respec¬
tively). The highest and lowest average uncorrected
pairwise distances and absolute genetic differences for
the 12S rRNA sequence alignment between the sportive
lemurs range from 9.6 to 0.7 and 75 to 9, respectively
( L . randrianasoli to L. jamesorum and L. mustelinus
to L. betsileo , respectively). The highest and lowest
average uncorrected pairwise distances and absolute
genetic differences for the PAST sequence alignment
between sportive lemurs range from 20.8 to 1.2 and
415 to 33, respectively (L. petteri to L.fleuratae and
L. ankaranensis to L. milanoii , respectively).
The phylogenetic inferences of the NJ and MP
analyses for the combined data sets for the genera
Avahi and Lepilemur are available as Appendices ll(s-
u). There is high bootstrap support for the MP and NJ
analyses with respect to the topology of the genera and
species. Appendices Ill(k-m) list the individual Avcihi
and Lepilemur haplotypes utilized in the various phylo¬
genetic inferences. By including GenBank accessioned
D-loop sequences and ND3 and ND4L sequences
(Figs. 5 and 6, respectively, and Appendix III(m)) from
Craul et al. (2007), L. otto was distinguishable from
the other 22 sportive species, but L. manasamody was
indistinguishable from L. grewcockorum. Additionally,
the Lepilemur species nova # 1 and Lepilemur species
nova #2, along with the sportive lemur from Mahilaka
(designated L. dorsalis ), were distinct from the other
sportive lemurs. The bootstrap support was high for the
interspecific relationships; however, the intraspecific
bootstrap was low. The minimum spanning network
presents diagrammatically the speciation among 24
sportive lemurs (Fig. 7).
20
Special Publications, Museum of Texas Tech University
IrU^lfr-niBr arrrJn
p™," . '..-m ■
m|
vplfcmur/tcmiViav
™ iL'frittmur iwifhtiit
.1LO-*
.tpfltmmr specie? rrmia
{T« ,‘i i HI J. F UDlf l'j
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I Ji'M'ii r jh (I r ra
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jijlgj l'i‘ | li. , i l rfflAir J.-iV. i^Iui
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TOP __
rJ'l.V'h Uf r.JH.Ai-i.Jrt J IIIIJI G (*. ,
Figure 4. Maximum parsimony phylogram derived from the PAST sequence data from
158 haplotypes from the 216 Lepilemur individuals with 19 out-group taxa (one of 5158
parsimonious trees). Values above branches indicate number of changes between nodes.
Values within circles along branches indicate support of bootstrap pseudoreplicates. Length
= 5158; Cl = 0.3806; RI = 0.9154; RC = 0.3484; HI = 0.6194. The subgroups of Lepilemur
species is as follows: A. northern sportive lemurs; B. southern sportive lemurs; C. west¬
ern sportive lemurs (west-central); D. eastern sportive lemurs; and E. western and eastern
sportive lemurs.
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
21
-A N P, ? ■ ■■' .ifit.n h ■i.rjb v .v
-RANOiSrt- Micmcvhits nifitx
ffiSSoW- Prcriimnrsitmis
—Huputemup aureus
I- Hapakmur griseus griseia
.'AR9 T -.l- Hapalemur griina acritfentaiu
-1-4S2- ■ • Varteia mrivgala wriegufa
kaNCms- ttatemur fiitna ntfus
11 -KANCUAS —-PmjNtfrecm edwarcisi
12 -MQR6*—- „ . —Prapithecw Wnwnm
MORSE
ImT -
-GaR£ -- Ckeirrtsithm media*
-HjVW>2?j- Cheirogak-us major
,RANC»7 I J T* . u
i| KA M)2iH F Avtiiu /.t i r lerasi
AhK, J3 -—-Arahi occidcfilalis
77 1
V
ZlOKOI .2
TYw i
1 .Un 1
•ty
kjRAj 1
MAI II
_ MAI 12 1
Lepi lemur edwardsi
Lepi lemur ahmansnnorum A
—Lepiiemur dorsalis
—Lepiiemur attkaraftensis
-Lepilemur milimoii
- L epifemu r t ym f tiacht sot 11
Lepiiemur petteri
Lepiiemur leucopus
- Lepiiemur mftcaudatus
. Lepiiemur randriartasoli
■Lepiiemur huhharrforum
-Lepiiemur aeeclis
Lepiiemur microdot i
Lepiiemur septen trsonaiis
—Lepiiemur sahamalensis
Lepiiemur otto
Lepiiemur gnewcockorum +
Lepiiemur weighing
-Lepiiemur betsileo
Lepiiemur mustelimts
Lepiiemur jamesorum
"• ■ ■■ EM U - Levttcmur jamesarun
- \we.i> l.epitemw flueretue
-A —,iaE2 . . Lepiiemur semi
Lepiiemur species nova #1
Lepiiemur species ttova #2
'• i m i Kt Ka
Figure 5. Maximum parsimony phylogram derived from the D-loop sequence data from 37 haplotypes
from the 69 Lepiiemur individuals with 19 out-group taxa (one of six most parsimonious trees). Val¬
ues above branches indicate number of changes between nodes. Values within circles along branches
indicate support of bootstrap pseudoreplicates. Length = 969; Cl = 0.4334; RI = 0.8057; RC = 0.3492;
HI = 0.5666. Accessioned sequences utilized from Craul et al. (2007). ^Designates GenBank ac¬
cession sequences for L. manasamody. A Designates the Lepiiemur dorsalis type at Antafondro and
Manongarivo. 'Designates the Lepiiemur dorsalis type at Mahilaka. GenBank accessions and base
size information is available in Appendix III(m).
22
Special Publications, Museum of Texas Tech University
- Cfcjmwleuf wtiitit
-kako:J *3 -- Chttwmkvt mafar
IKIAXI 24 I
-TrAN 033 -B|
—- TkA\iB<i r " f 'ityRkta ^ * »*TWF
•PntiittUif Mttinx
ANK 7 ——■■— Mfcw Mtm rtrVtftofcflftSri
RA ^OSiCl-ilj'cvTM'WjuY ntfiii
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ft A >*0332 ■—— PrupUhcvus afmittiri
MQS.6I1-_ - Propstb kus vtrrwmsxi
Emfl f— . A ptyrfmsi
ANK 33 *- AvahS tn-i-Hir/tlafix
•—■ Lepilemur edwardsi
Lepikmur grewoKkorum *
Lepilemur otto
J ..* Lepilemur microdon
BEZM Lepiiemur petteri
i"" ^'Lepilemur leucopus
■ yip jPljl .. Lepilem itr ruficaudatus
®Mt» . •Lepilemur hubbardorum
141 > ' 1 ^ .. 'Lepilem ur rtimlriumisofi
10 changes
Lepilemur ueeclis
Lepilemur ahmansonorum
'Lepilemur sahamtilensls
'"■Lepitemur aakaratutais
Lepilemur miianoii
Lepiiemur tymerlaehsom
Lepilemur dorsalis*
Lepilptntit tiarstili**
’Lepilemur septentrionalis
Lepilemur heistieo
•Lep Hem t ft jurn eso mm
■Lepiiemur mustelimis
Lepilemur jhietehie
- ’Lepilemur seall
.... . ■ ‘Lepilemur species nova #2
.i-.u aa a u *<Aasi 2 1 — 'Lepilemur species u<fm w l
' .... "Lepiiemur wiightae
Figure 6. Maximum parsimony phylogram derived from the ND3 and ND4L sequence data
from 41 haplotypes from the 69 Lepilemur individuals with 19 out-group taxa (one of 14 most
parsimonious trees). Values above branches indicate number of changes between nodes. Values
within circles along branches indicate support of bootstrap pseudoreplicates. Length = 1388; Cl
= 0.4135; RI = 0.8339; RC = 0.3449; HI = 0.5865. * Designates GenBank accession sequences
for L. manasamody. A Designates the Lepilemur dorsalis type at Antafondro and Manongarivo.
■Designates the Lepilemur dorsalis type at Mahilaka. Accessioned sequences utilized from Craul
et al. (2007). GenBank accessions and base size information is available in Appendix III(m).
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
23
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Special Publications, Museum of Texas Tech University
Discussion
General Observations and Considerations. —All
measurements were taken on live sedated animals,
which potentially produce inherent error from multiple
sources. Accurate age assessment by tooth size is often
a problem in cases where the individual is a subadult,
having not quite reached mature size, but no longer
an obvious juvenile. Since expeditions are carried
out in various habitats and forest fragments regardless
of weather and/or convenience, seasonal differences
can have an effect on the weights and measurements
based on food availability and reproductive status (e.g.,
pregnant or lactating). In food abundant seasons, some
anatomic reference points can be obscured by body con¬
dition. Furthermore, the accuracy of the morphological
data requires the same researcher take all measure¬
ments over the length of a study, which was logisti-
cally impossible. Additionally, a larger and equivalent
sample set is required to perform comprehensive and
thorough morphological analyses. Working with live
and subsequently released lemurs also precludes a
morphological comparison to museum type vouchers.
For these reasons, we interject the caveat to consider
the measurements as rough baselines that demonstrate
trends among the species of a genus providing the most
accurate morphometric data that could be collected
under field conditions.
Despite the limitations of the available morpho¬
logical data, some observations can be made from such
data. As Milne-Edwards (1875) suggested following
early explorations of Madagascar, substantiated by
Albrecht et al. (1990) and Godfrey et al. (1990), and
which Zaonarivelo et al. (2007) found in limited num¬
bers of Avahi species, we find the species inhabiting the
western forests are on average smaller than the species
inhabiting the east coast forests (Appendix ll(a-b); Ta¬
bles 2 A-C). However, this generalization is not evident
among the Lepilemur species (Appendix Il(c-d); Tables
3A-C). We find that L. leucopus and L. petteri (both
found on the southern tip of Madagascar) are on average
the smaller of the sportive lemur species described to
date. In addition, we determined that L. jamesorum ,
found in the Manombo Special Reserve, is more often
larger in measurements than other species.
Although the type locality of A. laniger is
considered to be Betanimena, Antongil Bay and Ma-
roanetsetra, this has not been conclusively determined
(Jenkins 1987; Groves 2001; Zaramody et al. 2006;
Andriantompohavana et al. 2007). As in Andriantom-
pohavana et al. (2007), the phenotypic and molecular
data (PAST fragment only ) reveal two types of A. la¬
niger , one type from Mantadia and another type from
Mananara-Nord National Park and Anjanaharibe-Sud
Special Reserve (Fig. 2; Appendix II(f, g, and i)). At
this point, the authors consider the taxonomy of A.
laniger unresolved at the species level. A comprehen¬
sive sample set incorporating the entire range from the
Mangoro River to Anjanaharibe-Sud Special Reserve,
along with representative data from the A. laniger Xy pe
specimen, needs to be evaluated in order to determine
species status and to assess whether the Onibe River
and/or Manigory River act as the limit or barrier for
both types or species.
Andriantompohavana et al. (2007) detected two
types of A. peyrierasi. In this study, our data present
three types of A. peyrierasi (Fig. 2; Appendix Il(e-i)).
All three types were sampled from forest fragments
south and north of Ranomafana National Park and all
types of A. peyrierasi can be found within the same
forest fragment. One interpretation of the current
sequence data is A. peyrierasi type #1, A. peyrierasi
type #3, and A. betsileo represent one species complex,
with A. peyrierasi #2 representing a distinct species
as A. peyrierasi. Again, results generated from larger
data sets over multiple generations, including pedigree
and karyotype data, will be required to resolve the
phylogenetic relationships between these three types
of A. peyrierasi.
The phylogenetic inference of the three mtDNA
regions, D-loop, 12S rRNA, and PAST fragments dif¬
ferentiated 23 species of sportive lemur, along with
detecting unrecognized biodiversity in the sportive
lemurs from Masoala and Mananara-Nord National
Parks, Lepilemur species nova # 1 and Lepilemur spe¬
cies nova #2 (Figs. 4-7; Appendices Il(j-r)). The data
corroborate the species status of L. otto in Craul et al.
(2007), but it did not distinguish L. manasamody from
L. grewcockorum (Figs. 5-6). The type localities of
both species are only two kilometers apart with no dis¬
cernible barrier (river or mountain; Zinner et al. 2007).
Even though a comparison of all sportive lemurs except
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
25
for L. mittermeieri was conducted in this study, only
partial GenBank sequences were available from other
investigations (for D-loop, Craul et al. (2007) accessed
386-388 bp while Louis et al. (2006b) accessed 534-
553 bp; for PAST, Craul etal. (2007) accessed 580-631
bp while Louis et al. (2006b) accessed 2359-2361 bp).
Therefore, in both fragments, sequence data had to be
truncated to enable comparative analyses, and this is
reflected in poor bootstrap support for interspecific as
opposed to intraspecific comparisons (Figs. 5-6; Ap¬
pendix III(m)). Since L. grewcockorum (Louis et al.
2006b) was published prior to Craul et al. (2007), L.
grewcockorum has priority over L. manasamody , the
junior synonym. The sportive lemur from Mananara-
Nord National Park, Lepilemur species nova #2, was
initially grouped within the species L. seali in Louis
et al. (2006b) due to the limited sample size. The ad¬
dition of the Masoala sportive lemurs to the phyloge¬
netic reconstruction and population aggregate analyses
distinguished this individual from the population of
sportive lemurs from Anjanaharibe-Sud Special Re¬
serve ( L. seali ) and Masoala National Park ( Lepilemur
species nova #1).
In Zinner et al. (2007), the authors reviewed
the current state of sportive lemurs of northwestern
Madagascar, highlighting the confusion over the type
locality of L. dorsalis and its generally accepted syn¬
onym, L. grandidieri. As shown in Figs. 5-6, multiple
distinct populations of sportive lemurs are designated
as L. dorsalis (Antafondro and Manongarivo refer to
L. dorsalis (2) in Zinner et al. (2007), and Mahilaka
individuals were not referenced in Zinner et al. (2007)).
Although the authors of this study agree with the con¬
clusions of Zinner et al. (2007) in that the holotypes of
both L. dorsalis and L. grandidieri need to be utilized
as comparative samples in molecular genetic analyses,
several points should be clarified. First, the sportive
lemurs from Antafondro and Manongarivo include
identical haplotypes even though they are found on
opposite sides of the Sambirano River, rather than
from the south side of the river as stated by Zinner
et al. (2007). Secondly, Zinner et al. (2007) did not
employ from all available investigations comparative
samples in their analyses, so not all of their conclusions
or interpretations can truly reflect the phylogenetic
relationships of the northwestern sportive lemurs (only
D-loop sequence data was evaluated and only a subset
from Louis et al. (2006b)).
Corroborating Evidence for Two New Lemur Spe¬
cies. —Multiple systematic studies utilizing molecular
genetic analyses, specifically mtDNA sequence and/or
cytogenetic data, have established this methodology to
be a reliable approach in distinguishing and character¬
izing unique diversity, and in this case, novel lemur
species (Rasoloarison et al. 2000; Andriaholinirina
et al. 2006; Andriantompohavana et al, 2006; Louis
et al. 2006a, 2006b; Rabarivola et al. 2006; Andrian¬
tompohavana et al. 2007; Olivieri et al. 2007). In this
study, we examined the taxonomic status of Avahi and
Lepilemur from Masoala National Park based on the
Phylogenetic Species Concept (Wheeler and Platnick
2000; Mayor et al. 2004; Louis et af 2006a, 2006b).
Since the authors consider the morphometrical data
as an insufficient sample set for the argument of es¬
tablishing species status, it has been provided as only
supplemental material. Although the morophological
data showed that multiple characters differed signifi¬
cantly between both geographically distant and adjacent
species ( Appendices ll(a-d); Tables 2C-3C), additional
samples would be necessary to apply a comprehensive
morphological analyses.
Three sets of criteria support the argument that the
two new putative species of lemurs (one Avahi and one
Lepilemur) should be considered distinct species. First,
the absolute genetic distances and average uncorrected
‘p’ distances between geographically adjacent species
and the proposed new species were within the limits
of other conspecifics (Appendix Ill(f-j); Olivieri et al.
2007). Secondly, the population aggregate analysis
(PAA) detected multiple diagnostic attributes for each
terminal clade (Tables 1 A-IE; Appendix Ill(a-e)). In¬
variably, as more individuals, populations, and species
are included in the data set, the number of diagnostic
attributes for each species will decrease due to random
mutations at previously unique base positions in even
more geographically distant species (Louis et al. 2006b;
Andriantompohavana et al. 2007; Olivieri et al. 2007).
Thirdly, these newly proposed species are located in
the relative isolation of the Masoala Peninsula where
other locally endemic taxa are to be found (e.g., Va-
recia rubra). Undoubtedly, additional morphometric
data from museum vouchers, nuclear DN A sequence
and genotype data, comprehensive samples from Ma-
nanara-Nord National Park, Anjanaharibe-Sud Special
Reserve, and surrounding forest fragments, along with
26
Special Publications, Museum of Texas Tech University
behavioral, ecological, and acoustical data should be
acquired to better characterize these new species.
Descriptive Diagnosis of Two New Species .—
The phylogenetic inference of the mtDNA sequence
data, along with the supplemental morphometric and
phenotypic character data, differentiate the described
species of woolly (genus Avahi) and sportive (genus
Lepilemur ) lemurs along with the previously unrec¬
ognized bioversity from Masoala National Park for
both genera, Avahi species nova #4 (NB # 1, #2, and #3
are discussed in Andriantompohavana et al. (2007))
and Lepilemur species novae HI and #2. Although the
sportive lemur from Mananara-Nord, Lepilemur species
nova #2, is considered distinct from other recognized
sportive lemurs, until additional samples are available
to confirm the reliability of the current information, the
Mananara-Nord sportive lemur will not be designated
as a proposed new species. A description of each new
species is provided below.
Avahi mooreorum , New Species
(formerly referred to as Avahi species nova #4)
Type Series. —Whole blood for MAS6.1
(TK145356; TTU-M 104519), adult female; MAS6.2
(TK145357; TTU-M 104520), adult male; and MAS6.3
(TK 145358; TTU-M 104521), adult male; are stored
and curated at Museum of Texas Tech University.
Individual measurements, e-voucher photos, and col¬
lection data are given in Avahi Field Data Appendix
1(a). MAS6.1, MAS6.2, and MAS6.3 were collected
by Edward Louis, Richard Rakotonomenjanahary,
Jean C. Randriamanana, Justin Andrianasolo, Philibert
Randriamaronjaka, and Jean Claude Rakotoniaina on
20 November 2006 and 21 November 2006.
Type Locality. —Madagascar: Province de
Antsiranana, Masoala National Park (approximately
S15°40'008", E049°57'476").
Description.—Avahi mooreorum (0.92 kg) is
slightly smaller than A. laniger (1.13 kg). The overall
pelage is a mixture (mottled) of chocolate brown and
light brown dorsum (Fig. 8). The ears are not readily
seen, blending in with the rest of the head. It has a
whitish patch under the mandible, and a distinct whit¬
ish patch, characteristic of other woolly lemurs, on the
posterior surface of the rear leg. A. mooreorum has a
diffuse pattern without the noticeable eyebrow of A. la-
niger. The facemask is apparent, but not as pronounced
as others, laniger (east coast) types (Andriantompoha¬
vana et al. 2007). The dorsum is darker in coloration on
the head as opposed to the distal portion of the dorsum
(gradual lighter pattern towards the tail base, which is
distinctly cream colored) The venter is gray including
undersides of limbs. The tail is reddish-brown with a
transition over the base to a cream color.
Diagnosis .—In the D-loop and PAST sequence
fragments, A. mooreorum differs from the closest rela¬
tive, A. laniger, by genetic and geographic distance
3.9%±0.8% (28 informative sites) and 4.2%±0.4% (104
informative sites), respectively. A. mooreorum has 28
diagnostic attributes (two attributes for D-loop and 26
attributes for PAST fragment).
Distribution. —Currently A. mooreorum is known
only from Masoala National Park. Even thought, la-
niger has been confirmed in Anjanaharibe-Sud Special
Reserve and Mananara-Nord National Park, further
surveys are required to confirm the northern extent of A.
mooreorum within Masoala National Park and possibly
intervening forest fragments between Anjanaharibe-
Sud Special Reserve and the Masoala Peninsula.
Comparisons and Remarks. — A. mooreorum
(0.92 kg) is smaller in weight than A. laniger (1.13
kg), however, A. mooreorum is slightly larger in body
length than A. laniger.
Etymology. —The name mooreorum is proposed
in honor of the Moore Family - Gordon and Betty
Moore, Ken and Kris Moore and Steve and Kathleen
Moore - in recognition of their long-term commitment
to biodiversity and conservation. The major support
that they have provided through the Gordon and Betty
Moore Family Foundation and other channels has
been critical to advancing conservation in some of the
world's most important and most endangered biodiv-
eristy hotspots, including Madagascar, and continues
to have enormous impact.
Vernacular Names. —Moore’s woolly lemur or
Masoala woolly lemur.
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
27
Figure 8. Avahi mooreorum , Moore’s woolly lemur. Photo by Edward E. Louis, Jr.
28
Special Publications, Museum of Texas Tech University
Lepilemur scottorum , New Species
(formerly referred to as Lepilemur species nova #1)
Type Series. —Whole blood for MAS6.12
(TK145333; TTU-M 104516), adult female; MAS6.13
(TK145334; TTU-M 104517), adult female; and
MAS6.14 (TK145335; TTU-M 104518), adult male;
are stored and curated at the Museum of Texas Tech
University. Individual measurements, e-voucher
photos, and collection data are given in Lepilemur
Field Data Appendix 1(a). MAS6.12, MAS6.13 and
MAS6.14 were collected by Edward Louis, Richard
Rakotonomenjanahary, JeanC. Randriamanana, Justin
Andrianasolo, Philibert Randriamaronjaka, and Jean
Claude Rakotoniaina on 20 November 2006.
Type Locality. —Madagascar; Province de
Antsiranana, Masoala National Park (approximately
S15°40'246", E049°57'537").
Description.—Lepilemur scottorum is a me¬
dium-sized sportive lemur (0.88 kg) with a uniform
reddish-brown color pattern (Fig. 9). The venter is
reddish brown. The pelage is long and thick through¬
out the body, with a diffuse black stripe, midline of the
dorsum that ends mid-body. The face is whitish-gray.
The cheeks and eyebrows are white. The hands and
feet are reddish brown. The tail is reddish brown at
the base and progressively brownish-gray towards the
tip. The tip is black in coloration.
Diagnosis. —In the D-loop, 12S rRNA, and
PAST sequence fragments, L. scottorum differs from
its closest relatives, L. seali and L. species nova# 2, by
genetic and geographic distance 6.3%±1.1% (36 infor¬
mative sites) and 7.0%±1.2% (37 informative sites),
2.2%±0.5% (18 informative sites) and 1,8%±0.5% (15
informative sites), and 5.1%±0.5% (116 informative
sites) and 4.2%±0.4% (95 informative sites), respec¬
tively. L. scottorum has 16 diagnostic attributes (six
attributes for D-loop, 2 attributes for 12S rRNA, and
8 attributes for PAST fragment).
Distribution. —Currently known only from
Masoala National Park, Masiaposa Forest.
Comparisons and Remarks.—Lepilemur scot¬
torum (0.88 kg) is smaller than L. seali (0.96 kg), L.
mustelinus (0.99 kg), and L. wrightae (1.15 kg). Addi¬
tional survey work is required to determine the eastern
and northern range of L. scottorum.
Etymology. —The name scottorum is proposed
in honor of the Suzanne and Walter Scott Jr. Fam¬
ily Foundation. Suzanne and Walter Scott Jr. are
definitive leaders, mentors, and motivators of in situ
and ex situ conservation throughout the world and
have volunteered extensively in Henry Doorly Zoo’s
conservation programs for decades. Walter Scott, Jr.
is a 32-year member of the Board of Directors of the
Omaha Zoological Society. Formerly president of the
board, he has been Chairman of the board since 1984.
He is a philanthropist, business leader, and community
activist on the board of many charitable organizations
and businesses. Suzanne M. Scott, a long-time zoo
supporter, in 1984 became the founding executive
director of the Omaha Zoo Foundation responsible for
establishing the zoo’s endowment funds and raising
money for special projects.
Vernacular Names. —Scott’s sportive lemur.
In conclusion, Madagscar has lost approximately
90% of its original forest due to human activities
such as agriculture and logging. Exacerbating this
loss, the remaining forests have become increasingly
isolated, resulting in an unconnected mosaic of forest
fragments (Ganzhorn et al. 2001). The extreme levels
of deforestation have left many species susceptible to
stochastic events or anthropogenic pressures (Jernvall
and Wright 1998; Ezard and Travis 2006). Moreover,
there is insufficient information available on distribu¬
tions of described lemurs across Madagascar. The
current distributions of the genera Avahi (nine species)
and Lepilemur (24 species) are presented in Figs. 10
and 11, respectively. The impact of rivers as barriers
and the remarkable influence of isolation on allopatric
speciation is diagrammatically demonstrated in Fig.
12. The inter-river system (IRS) for the genera Mi-
crocebus and Lepilemur in northwestern Madagascar
as described by Olivieri et al. (2007) and Craul et al.
(2007), respectively, can be vividly expanded across
the island as unique paired biodiversity exists for the
genera Avahi and Lepilemur wherever they are found.
It is urgent to identify all biodiversity and species’
distributions before the 2008 commitment to triple
the amount of protected area by the Malagasy govern¬
ment. This data can only enhance the delineation of
the reserve matrix by providing valuable information
for wildlife and conservation agencies and allowing
precise management decisions to be implemented to
protect the existing biodiversity.
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
29
Figure 9. Lepilemur scottorum, Scott’s sportive lemur. Photo by Edward E. Louis, Jr.
30
Special Publications, Museum of Texas Tech University
Bcmarivo
Maningory
“~~~GnLbe
Mangold
Faraony
Mahavavy Nord-
Antafondro (Maromiandra)'
Sambirnrio-
A 0 J 00
km
Mahajamba
Mariarano
Beisiboka
Ankarafantsika
Mahavavy Sud
Maningoza
Manambaho
Manambob
Irodo
Laky
Tsingy de
Bcmaraha
Tsiribihina
Morondava
Fibers liana
Onilahy
Maevarano
Sofia-—
Lwtfl
Mcrurundnj
Anjanaharibe-Siid
Antainambalana
Masoala (Masiaposa)
Mananara-Nord (I von tak a-S ud)
Mananara-Nord (Verezananatsoro)
Mantadia
Maromizaha
Mangoro
Fandriana
Nanfiomna
Ranomafana
Manampacrana
Manombo
Mananara
Andohahela
(Manangoiry)
Mandrarc
UP Avaki betsltto
^ 4 vtifti mpareariim
^ .4 va h i p eyri eras i
^ .1 vahi rutiumsintxttiivana
^ Avaki mtrtdtonails
^ Avahi lauiger
^ Av<ihi tuticolor
A vahi ctetsei
I Avahi accidental!*
^ Emil CO tor Luf site denirtcH
west coasl Avafti species,
all formerly Avahi aciidenlalis
4) Fori I color of site denotes
cast coast Avahi species,
all formerly 4 vb/jj lumber
Figure 10. Revised distribution map of the woolly lemurs (genus Avahi) of Madagascar.
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
31
It
5.
S' •
|
ill
* S S
l!
I I
It
^ =i.
1 a ! i 11
alls gai
a § S 1 g 1 f
£.S,-S
t % k s s s I
mini
t&ttttt
o r *J ^ -J -J -J
^ • ••
s
. 1
la J
"IU
z i s s s s s s
| 6 B 8 s ! 5
jtttttt
>^>hl ^ -4 ^
CJ 2 -s
~ 3
Figure 11. Revised distribution map of the sportive lemurs (genus Lepilemur ) of Madagascar.
32
Special Publications, Museum of Texas Tech University
Figure 12. Combined distribution map of the genera Avahi and Lepilemur. A minimum of one Avahi species and
one Lepilemur species are located between river pairs that act as barriers or boundaries of the range. Rivers that
segregate Avahi and Lepilemur species on the western half of Madagascar are displayed in red font, and the rivers
that segregate Avahi and Lepilemur species on the eastern half of Madagascar are in blue font.
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
33
Acknowledgments
This manuscript was supported by a grant from
the Primate Action Fund and Conservation Interna¬
tional, Inc. This project would not have been possible
without the support of the staff, guides, and drivers
of the Institute for Conservation of Tropical Environ¬
ments, Madagascar (ICTE-M1CET), as well as the
Association Nationale pour la Gestion des Aires Pro¬
tegees (ANGAP), Parc Botanique et Zoologique de
Tsimbazaza, U.S. Fish & Wildlife Service, University
of Antananarivo’s Anthropology and Paleontology
Department, and the Ministere des Eaux et Forets of
Madagascar. We acknowledge the generosity of Bill
and Berniece Grewcock for their long-term support and
commitment, which gave the Center for Conservation
and Research (Henry Doorly Zoo) its direction and
identity. Furthermore, we would like to acknowledge
that this research would not be possible without the
incredible support of the Ahmanson Foundation, the
James Family Foundation, and the Theodore F. and
Claire M. Hubbard Family Foundation. Although
listed together, this does not diminish each founda¬
tions contribution to the Malagasy graduate students
and held assistants that their long-term contributions
have sustained. We also would like to acknowledge
the graphics specialist, Lisa Reinhart, for creating the
web page documents.
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Addresses of authors:
Runhua Lei
Center for Conservation and Research
He my Doorly Zoo
370iS. 10 1h St.
Omaha, NE 68107, USA
E-mail: leirfajomahazoo. com
Shannon E. Engberg
Center for Conservation and Research
Henry Doorly Zoo
3701S. 10 th St.
Omaha, NE 68107, USA
E-mail: 2 enetics(a>omahazoo. com
Rambinintsoa Andriantompohavana
University of Antananarivo
BP 906
Antananarivo 101, Madagascar
E-mail: radambvHDZ(d)vahoo. fr
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population of Sambirano woolly lemur ( Avahi unicolor)
from the Anaborano Forest in northwestern Madagascar.
Lemur News 12:21-24
Zaramody, A., J.-L. Fausser, C. Roos, D. Zinner, N. Andriaho-
linirina, C. Rabarivola, I. Norscia, I. Tattersall, and Y.
Rumpler. 2006. Molecular phylogeny and taxonomic
revision of the eastern woolly lemur {Avahi laniger).
Primate Report 74:9-22.
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challenges for mitochondrial DNA markers. Trends in
Ecology and Evolution 11:247-251.
Zinner, D., C. Roos, J.-L. Fausser, C. Groves, and Y. Rumpler.
2007. Disputed taxonomy classification of sportive
lemurs {Lepiiemur) in NW Madagasacar. Lemur News
12:53-56.
Susie M. McGuire
Center for Conservation and Research
Henry Doorly Zoo
3701 S. 10 th St.
Omaha, NE 68107, USA
E-mail: genetics(a).omahazoo. com
Russell A. Mittermeier
Conservation International
2011 Ctystal Drive Suite 500
Arlington, VA 22202, USA
E-mail: t: mittermeierfa).conservation, org
John R. Zaonarivelo
University of Antananarivo
BP 906 '
Antananarivo 101, Madagascar
E-mail: zaonarivelo(d)vahoo. fr
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
37
Addresses of authors (cont.):
Rick A. Brenneman
Center for Conservation and Research
Heniy Doorly Zoo
3701 S. 10 th St.
Omaha, NE 68107, USA
E-mail: rabr(a),omahazoo. com
Edward E. Louis, Jr.
Center for Conservation and Research
Henry Doorly Zoo
3701S. 10 ,h St.
Omaha, NE 68107, USA
E-mail: edlo(a),omahazoo. com
E-mail: (Lab - United States) :
senetics(a),omahazoo. com
Email: (Field - Madagascar):
kelvnewsl(a),vahoo. com
The following Appendices I-III to this publication are available online at the indicated website addresses.
Appendix I
a. Avahi and Lepilemur Field Data Appendix (Individual data file for each newly added Avahi and Lepilemur ,
including morphometries, photos, sequence accessions, global position system, microchip data, gender, and
location).
http://10.1Q.10.3/ccr/genetics/lemur/index.asp?page=ccr/genetics/lemur/^vaIgp/MAS.htm
b. Avahi and Lepilemur table of individual samples and corresponding information for each sample (bar code
number, site, original species designation, current species designation, GenBank accession numbers of sequence
data).
http://www.omahazoo.com/ccr/genetics/papers/ Appendix\AvahiLepilemurTab\Q.pdf
38
Special Publications, Museum of Texas Tech University
Appendix II
a. Appendix II histograms of means and 95% confidence intervals for significant morphometric parameters found
among 11 Avahi species or types.
http://www.omahazoo.eom/ccr/genetics/papers/AppendixIL4va/7/HistogramPartI.pdf
b. Appendix II histograms of means and 95% confidence intervals for significant morphometric parameters found
among 11 Avahi species or types.
http://www.omahazoo.com/ccr/genetics/papers/AppendixIL4va/77HistogramPartII.pdf
c. Appendix II morphometric parameter averages and standard deviations among 19 Lepilemur species.
http://www.omahazoo.com/ccr/genetics/papers/AppendixILLgp/HistogramPartl.pdf
d. Appendix II morphometric parameter averages and standard deviations among 19 Lepilemur species.
http://www.omahazoo.com/ccr/genetics/papers/AppendixHXgp/HistogramPartILpdf
e. Appendix II Avahi D-loop fragment haplotypes maximum likelihood phylogram.
http://www.omahazoo.eom/ccr/genetics/papers/appendixIL4va/7/DLPHaploML.pdf
f. Appendix II Avahi PAST fragment haplotypes maximum parsimony phylogram.
http: //www. omahazoo. com/ccr/genetics/papers/AppendixIL4va/?/PASTHaploMP pdf
g. Appendix II Avahi PAST fragment haplotype maximum likelihood phylogram.
http ://www. omahazoo, com/ccr/genetics/papers/AppendixIL4va/?/PASTHaploML ,pdf
h. Appendix II Avahi D-loop fragment haplotype Bayesian analysis cladogram.
http://www.omahazoo.eom/ccr/genetics/papers/AppendixIL4va/7/DLPHaploBavesian.pdf
i. Appendix II Avahi PAST fragment haplotype Bayesian analysis cladogram.
http://www.omahazoo.eom/ccr/genetics/papers/AppendixIL4va/7/PASTHapBavesian.pdf
j. Appendix II Lepilemur D-loop fragment haplotype maximum parsimony phylogram.
http://www.omahazoo.com/ccr/genetics/papers/AppendixIIXgp/7g7777/7DLPHaploMP.pdf
k. Appendix II Lepilemur D-loop fragment haplotype maximum likelihood phylogram.
http://www. omahazoo.com/ccr/genetics/papers/AppendixIIXgp//g7777//^DLPHaploML.pdf
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
39
Appendix II (cont.)
l. Appendix II Lepilemur D-loop fragment haplotype Bayesian analysis cladogram.
http://www.omahazoo.eom/ccr/genetics/papers/AppendixIILe/7/yDLPHapBavesian.pdf
m. Appendix II Lepilemur PAST fragment haplotype maximum likelihood phylogram.
http: //www. omahazoo, com/ccr/genetics/papers/AppendixIILep/PASTHaploML .pdf
n. Appendix II Lepilemur PAST fragment haplotype Bayesian analysis cladogram.
http://www.omahazoo.com/ccr/genetics/papers/AppendixIILez7zPASTHaploBavesian.pdf
o. Appendix II Lepilemur 12S rRNA fragment haplotype neighbor-joining phylogram.
http://www.omahazoo.eom/ccr/genetics/papers/AppendLxIILe/7/T2SrRNAHapNJ.pdf
p. Appendix II Lepilemur 12S rRNA fragment haplotype maximum parsimony phylogram.
http://www. omahazoo.com/ccr/genetics/papers/Append ixIILe/7/12 SrRNAHapMP.pdf
q. Appendix II Lepilemur 12S rRNA fragment haplotype maximum likelihood phylogram.
http: //www. omahazoo. com/ccr/genetics/papers/AppendixIILe/7/12 SrRNAHapML, pdf
r. Appendix II Lepilemur 12S rRNA fragment haplotype Bayesian analysis cladogram.
http ://www. omahazoo, com/ccr/genetics/papers/AppendixIILe/7/12 SHapBavesian.pdf
s. Appendix II Lepilemur and Avahi neighbor-joining phylogram derived from the PAST fragment sequence data
from 119 Avahi individuals and 216 Lepilemur individuals with 25 out-group taxa. Species designated according
to distribution in the current literature (Thalmann and Geissmann 2000,2005; Andriaholinirina et al. 2006; Louis
et al. 2006b; Mittermeier et al. 2006; Zaramody et al. 2006; Andriantompohavana et al. 2007).
http://www.omahazoo,com/ccr/genetics/papers/AppendixIILc/7/Tvcz/?z PASTHaploNJ.pdf
t. Appendix II Lepilemur and Avahi neighbor-joining phylogram derived from the D-loop fragment sequence data
from 119 Avahi individuals and 216 Lepilemur individuals with 25 out-group taxa. Species designated according
to distribution in the current literature.
http ://www. omahazoo, corrPccr/genetics/papers/AppendixLeff/H iy/L/ DLPHaploNJ. pdf
u. Appendix II Avahi and Lepilemur D-loop and PAST combined fragment haplotype maximum parsimony
phylogram.
http://www.omahazoo.eom/ccr/genetics/papers/appendhLJwz/ 7 /Lgz 7 z/e/ 72 //rHapMP.pdf
40
Special Publications, Museum of Texas Tech University
Appendix III
a. Table 1A. Diagnostic nucleotide sites from the D-loop fragment Pairwise Aggregate Analysis (PAA) of
Avahi.
http://www.omahazoo.eom/ccr/genetics/papers/appendixIILdvf3/7/DLPPAA.pdf
b. Table IB. Diagnostic nucleotide sites from the PAST fragment Pairwise Aggregate Analysis (PAA) of
Avahi.
http://www.omahazoo.eom/ccr/genetics/papers/appendixIIL4vf3///PASTPAA.pdf
c. Table 1C. Diagnostic nucleotide sites from the D-loop fragment Pairwise Aggregate Analysis (PAA) for
Lepilemur.
http://www.omahazoo.eom/ccr/genetics/papers/appendixIILLe/7/DLPPAA.pdf
d. Table ID. Diagnostic nucleotide sites from the 12S rRNA fragment Pairwise Aggregate Analysis (PAA) of
Lepilemur.
http: //www. omahazoo, com/ccr/genetic s/papers/ appendixIIIZc/7/ 12 S rRNAPAA.pdf
e. Table IE. Diagnostic nucleotide sites from the PAST fragment Pairwise Aggregate Analysis (PAA) of Lep¬
ilemur.
http: //www. omahazoo, com/ccr/genetic s/papers/ appendixIIIZc/7/PAS TPAA, pdf
f. Table 5A. Genetic distance matrix for D-Loop fragment sequence data for Avahi species.
http://www.omahazoo.eom/ccr/geneties/papers/appendixIIL4vf3/7/DLPGDM.pdf
g. Table 5B. Genetic distance matrix for PAST fragment sequence data for Avahi species.
http://www.omahazoo.eom/ccr/geneties/papers/appendixIIL4vf3///PASTGDM.pdf
h. Table 6A. Genetic distance matrix for D-Loop fragment sequence data for Lepilemur species.
http://www.omahazoo.com/ccr/genetics/papers/appendixIIIZc/7zDLPGDM.pdf
i. Table 6B. Genetic distance matrix for 12S rRNA fragment sequence data for Lepilemur species.
http ://www. omahazoo. com/ccr/genetics/papers/appendixIIIZc/7z 12 SrRNAGDM. pdf
j. Table 6C. Genetic distance matrix for PAST fragment sequence data for Lepilemur species.
http://www. omahazoo. com/ccr/genetics/papers/appendixIIIZc/7/PASTGDM.pdf
Lei et al.— Nocturnal Lemur Diversity at Masoala National Park
41
Appendix III (cont.)
k. Appendix III Haplotype Table I (Summary of designated haplotypes for the genus Avahi from all localities
for D-loop and PAST fragments).
http://www.omahazoo.eom/ccr/genetics/papers/AppendixIILAY//7/HaploWpeTableI.pdf
l. Appendix III Haplotype Table II (Summary of designated haplotypes for the genus Lepilemur from all localities
for D-loop, 12S rRNA, and PAST fragments).
http://www.omahazoo.com/ccr/genetics/papers/AppendixIIILepilemurHapTableII.pdf
m. Appendix III Haplotype Table III ( Summary of designated haplotypes for the genus Lepilemur from localities
for D-loop, ND3 and ND4L fragments from Craul et al. (2007).
http://www.omahazoo.com/ccr/genetics/papers/AppendixIIILepilemurCraulTableIII.pdf
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